animals

Article
Food Preferences in Cats: Effect of Dietary
Composition and Intrinsic Variables on Diet Selection
Raúl A. Alegría-Morán 1,2 , Sergio A. Guzmán-Pino 3 , Juan I. Egaña 3 , Valeria Sotomayor 3
and Jaime Figueroa 4, *
1 Departamento de Medicina Preventiva Animal, Facultad de Ciencias Veterinarias y Pecuarias,
Universidad de Chile, Santiago 8820808, Chile; ralegria@veterinaria.uchile.cl
2 Facultad de Ciencias Agropecuarias, Universidad Pedro de Valdivia, Santiago 7500908, Chile
3 Departamento de Fomento de la Producción Animal, Facultad de Ciencias Veterinarias y Pecuarias,
Universidad de Chile, Santa Rosa 11735, La Pintana, Santiago 8820000, Chile;
sguzmanp@uchile.cl (S.A.G.-P.); jegana@uchile.cl (J.I.E.); valeriasotomayor.l@gmail.com (V.S.)
4 Departamento de Ciencias Animales, Facultad de Agronomía e Ingeniería Forestal,
Pontificia Universidad Católica de Chile, Santiago 7820436, Chile
* Correspondence: figueroa.jaime@uc.cl; Tel.: +56-22-3544092




Received: 13 April 2019; Accepted: 17 June 2019; Published: 19 June 2019


Simple Summary: Cats tend to retain food preferences across their lifetime, but intrinsic variables
could affect their diet choices so as to preserve internal homoeostasis. This work intended to study
the feeding behavior of cats in relation to diet composition and some intrinsic variables (sex, age,
and body weight) by analyzing data from a 10-year database of two-feeder food preference tests
(2007–2017). Diet mineral components (calcium, crude fiber and ashes) affected food preferences
negatively. In addition, the influence of body weight and sex manifested as lower food intake in both
females and the heaviest cats (relative to bodyweight). However, only body weight affected food
preferences, where the heaviest animals presented the higher preferences overall. During the cold
season, animals (especially females) displayed higher food intake, whereas hot seasons increased
male cat preferences for palatable diets, hence we observed an interaction between sex and climate
season. In conclusion, understanding the relationship that both food composition and some intrinsic
variables cats have with their diet preference could help in improving the formulation of specific pet
food diets, so that these adequately satisfy the physiological and hedonic needs of domestic cats.

Abstract: A ten-year database of food preference tests (n = 1021; period 2007−2017) was used
to explore the feeding behavior of domestic cats. Principal component (PC) analysis and linear
regression between food nutrients and preferences (for the most preferred diet of each test; Diet A)
were performed. Intake and preference for Diet A were analyzed by intrinsic cats’ variables and
climate season. The PC1 (calcium (Ca), phosphorus (P), and ash), PC2 (lipids and ether extract)
and PC4 (crude fiber; CF) had borderline significance (p < 0.06; β = −1.42, β = −1.56, and β = 2.68,
respectively). Ash and CF contents presented negative correlations with food preference (rho = −0.269,
p = 0.031; rho = −0.338, p = 0.006, respectively), and Ca had borderline significance and negative
correlation with food preference (rho = −0.241, p = 0.054). Body weight and sex influenced the intake
of Diet A, being lower for females (β = 11.758; p = 0.014) and heaviest cats (β = −5.490; p < 0.001).
However, only body weight affected food preferences, where the heaviest cats had greater preferences
for Diet A. Hot season decreased food intake (β = −2,117; p = 0.032), mostly in females (rho = −3.537;
p = 0.002). Males had greater preferences for Diet A during hot seasons (β = 10.216; p = 0.023) and
females presented similar preferences throughout the year (p = 0.950). Mineral contents, body weight
and sex affected food intake and preferences of cats under the influence of climate season, probably
explained by adaptive changes in food detection.

Keywords: cat; commercial diets; food selection; nutrients; climate season; sex

Animals 2019, 9, 372; doi:10.3390/ani9060372 www.mdpi.com/journal/animals

Animals 2019, 9, 372 2 of 13

1. Introduction
Domestic cats (Felis silvestris catus) display feeding behavior that stems from specific nutritional
requirements [1–3] associated to their strict carnivore physiology, such as high levels of total crude
protein linked to the presence of specific amino acids in the diet (i.e., arginine and taurine), vitamins
A, D, and complex B, as well as arachidonic acid [4–6]. Usually, cats eat small portions throughout
the day mimicking a feeding rhythm pattern that is typical of their wild cat ancestors (Felis silvestris
lybica), who hunted small-sized prey [5,7]. Cats choose their diets based on smell, taste, temperature,
and texture [6,8,9] up to the point of self-regulating consumption of certain kinds of foods to ensure
an adequate intake of certain nutrients, hence balancing their diets themselves [10,11]. In this regard,
cats are innately drawn to foods with a strong umami flavor, which is typically linked to a high
concentration of amino acids [5].
Usually, cats show little interest in eating foods that are rich in sugar and some studies have
confirmed that they do not perceive the sweet taste [5]. Hence, cats forgo sweet foods and limit their
energy intake to less than 300 kJ/day, and in case of consuming high amounts of these nutrients,
they typically vomit and display diarrhea afterwards [12]. Similarly, domestic cats and their ancestors
had a much higher threshold (>0.05 M) than other species in regard to salt in food, which seems to fit
with their nutritional requirements of sodium chloride that are covered by the high content of salt
present in their regular carnivore diet [4,5,12]. Cats can react differently to new ingredients in their diet
depending on their origin, experience, and nutritional status, ranging from caution and neophobia
so to avoid possible intoxication or harm [13], to neophilia that results in greater intake of new and
less readily available diets [14,15]. The latter situation has been observed in both wild and farm cats,
helping them to broaden their food variability and meet their total nutritional requirements [14,15].
The pet industry constantly performs tests to assess food preferences in cats with the intention
of understanding the variability of their choices when offered a wide range of flavors, ingredients
or presentations. Furthermore, the industry is also trying to offer a range of food products catering
to nutritional and hedonic requirements of cats according to intrinsic variables, such as age, breed,
reproductive status, weight, or even specific health conditions [5]. However, few scientific studies have
explored how intrinsic variables might alter the magnitude of preferences for different commercial cat
diets [5]. Cats usually are able to control their daily calorie intake even if they are offered free access to
commercial diets [16,17]. Nevertheless, obese cats cannot self-regulate their calorie intake, displaying
changes in their feeding patterns that are probably due to an altered perception of macronutrients,
as well as a deregulation of other physiological ingestion mechanisms that have been described for
humans [6]. In the case of humans, their tolerance and preference for high-fat foods is influenced by the
fact that people with a normal body mass index (BMI) have a lower threshold for detecting lipids than
people whose BMI is higher [18]. When considering other variables, such as sex, researchers have found
that males present lower thresholds for hunting behaviors than females, and when males are neutered,
they increase their latency for attacks [19]. Meanwhile, both cats and humans experience physiological
changes in regard to their feeding behavior that stem from their age, such as a decreased digestive
capacity, less motor activity, and lower metabolic requirements [4,6]. Age can also affect reward
areas in the brain, resulting in diet acceptance differences due to individual changes in palatability
perception [20,21]. Similarly, age could also impact on olfactory and gustatory systems that trigger food
preference changes and cautious feeding behavior in cats [22]. Given this background, the current study
intends to explore how the preference for commercial diets could be explained by either nutritional
components of the diets or by intrinsic variables of cats.
Animals 2019, 9, 372 3 of 13

2. Materials and Methods

A database covering 10 years of cat food preferences (2007−2017) was provided by the Research
Centre for Pet Feeding Behaviour (FAVET–Faculty of Veterinary and Animal Sciences, University
of Chile), located in the metropolitan region of Chile (Santiago city, 34◦ 210 S, 71◦ 180 W), and then
used to analyze the effect of diet composition and intrinsic variables of cats on their food intake and
preference for commercial diets. Experimental procedures were approved by the bioethical committee
of “Facultad de Ciencias Veterinarias y Pecuarias”, Universidad de Chile (N◦ 042013).

2.1. Animals and Housing

A total of 1021 preference tests using 24 different mongrel domestic cats (Felis silvestris catus)
(9 male and 15 female) were used in this study over the complete period (2007–2017). An average of
15 ± 2 animals was maintained per year at the Center. These animals were sourced from local shelters
at an average age of 6 months and then submitted for clinical examination by veterinarians of the
Clinical Sciences Department of FAVET before being registered as experimental animals. Once housed,
they were trained for 3 to 6 months to ease their adaptation to the experimental procedures. During this
period, they were adapted to 2-feeder preference trials with highly palatable foods in both feeders to
stimulate the intake of the two options. Health of cats was assessed prior to the start of the experiments
through clinical, hematologic and biochemical serum profiles, and all cats that participated in preference
trials were considered healthy. The age of the cats ranged from 1 to 13 years old and females usually
represented a greater percentage of the total animals. Cats were housed individually in single kennels
that were split into inner and outer areas of 94 × 90 × 105 cm and 94 × 90 × 90 cm, respectively.
Cats had access to a sandbox and fresh water on the outer area of each kennel, while on the
inner area animals were fed a single ration (3% of their body weight) of a commercial diet (daily at
09:30 a.m.), as well as ad libitum access to water and a comfortable bed. This ration was adjusted
monthly accordingly to body weight records of each cat to ensure their body condition remained stable.
Environmental conditions were partially controlled by the Centre by means of door and window
management. However, average temperature varied across different seasons during the year, ranging
between a maximum of 22 ◦ C in summer and a minimum of 7.7 ◦ C in winter [23]. Animals had access
to a common playground area (300 × 375 × 480 cm) for 1 h each day, provided with environmental
enrichment objects, where they could exercise. Additionally, veterinary staff from FAVET regularly
checked the animals and performed clinical examinations to assess their health status. In general,
cats ended their participation in experimental trials when they reached 8 years of age (with some
exceptions) or when they presented diseases not compatible with the preference tests or that could
alter their results.

2.2. Procedures to Determine Food Preferences and Database

Food trials comprised a series of two-bowl food preference tests that were performed within each
kennel to assess the variability of their nutritional, physical, and/or organoleptic properties. Each trial
lasted 4 days (1 test/day) in which diets were counterbalanced in position (left or right) to diminish
the effect of possible side bias over preferences. Every test began on the afternoon (02:00 pm) and
ended on the next morning (10:00 am). Each cat was individually tested over this 20-hour period by
offering two different diets, each on one bowl of the same color, shape, and size. Rations offered were
adjusted according to their body weight (3%) at all times, whether animals were participating in a trial
or not. On test days, cats were fed only the experimental diets and water was freely available for them.
Feeders were weighed at the beginning and the end of each test, to estimate the amount of food that
the animals consumed from each diet. In order to better equate intake across animals and diminish size
effects, consumption was subsequently corrected by their metabolic weight (total consumption/body
weight0.75 ). As for the nutritional composition of the experimental diets, these were subjected to a
proximate chemical analysis, acid hydrolysis, and a calorimetric pump (IKA® calorimetric system C
Animals 2019, 9, 372 4 of 13

2000, Wilmington, NC, USA). Diets were analyzed following standard Association of Official Analytical
Chemist (AOAC) methods [24] for content of moisture (method 945.15), crude protein (CP, Kjeldahl
method 945.18, N × 6.25), ether extract (EE, method 945.16), ash (method 920.153), crude fiber (CF,
method 962.09), total lipids (LIP, method 922.06), calcium (Ca), phosphorus (P), dry matter (DM),
nitrogen free extract (NFE) and metabolizable energy (ME).
With the objective to homogenize and compare different studies performed across the whole
period (2007–2017) without having the same standard diet across tests, the most preferred diet in
each test was labeled as Diet A, whereas the less preferred one was labeled as Diet B, and the degree
of preference was expressed as the relative consumption of Diet A over Diet B + Diet A allowing
evaluation of the effect of variables on the selection of the most preferred diet (see Equation (1)). To this
end, our data for consumption and preference of Diet A were sorted by sex, age, and body weight of
each cat, as well as the current climate season at the time of performing each test.

Intake of Diet A
× 100 (1)
Intake of Diet A + Intake of Diet B

2.3. Statistical Analysis

2.3.1. Effect of Nutrient Composition over Cats’ Food Preferences

The evaluation of how nutritional components of diets (DM, CP, CF, EE, NFE, ash, Ca, P, LIP
and ME) may explain food preferences and if the components were grouped, was carried out using a
principal component analysis (PCA) and ‘devtools’ and ‘ggbiplot’ packages of the statistical software
R [25]. All nutritional components were expressed as the net difference between Diet A and B (only
metabolic energy was expressed as the percentage difference between diets). After the PCA and
multivariable linear regression was performed confronting the most important variables grouped in
the PCA and cats’ preferences by using R statistic software. Spearman correlations were performed
with the nutritional elements that represented the greatest variability within the significant principal
component (PC) obtained from the prior analysis.

2.3.2. Effect of Intrinsic Variables Over Cats’ Food Preferences

Intrinsic variables effects on intake and preference of Diet A was examined using linear mixed
models, also known as variance component models, which are characterized by the consideration
of the nested structure of the data, providing a chance to decompose variance into a number of
different components which can then be given a useful interpretation [26–28]. The intrinsic variables
recorded in this study correspond to one categorical variable, sex (male or female); body weight
was registered as a continuous variable. Season was also recorded, considering cold season as all
the experiments performed between March 21 and September 20 (autumn and winter seasons in the
southern hemisphere) and hot season as all the experiments performed between September 21 and
March 20 (spring and summer seasons in the southern hemisphere); this variable was also recorded as
each one of the traditional seasons (summer, autumn, winter, and spring) according to the southern
hemisphere dates. Two dependent variables were analyzed (intake and food preference), leading to the
generation of two models to determine the effect of the registered variables over intake and preference
of Diet A in cats. Cat variable was treated as a random effect, while the rest of the variables were
treated as fixed effects. The proposed model to explain intake and preference is:

Ykm = β0 + βn + bk + εm(k) (2)

where Ykm corresponds to the intake (corrected for metabolic weight) or to Diet A preference, β0 is
the intercept, βn is the vector of fixed effects, bk corresponds to the random effect of the kth cat and
εm(k) is the error term or the elements that were not included in the analysis. This model assumes that
random effects and errors are independent at the nesting level and follow an approximately normal
Animals 2019, 9, 372 5 of 13

distribution, with means equal to zero and variance equal to σ2 . The Likelihood Ratio Test (LRT) was
used to select fixed effects from the initial model (full model), removing variables whose regression
coefficients were not significant (p > 0.05) using a stepwise backward elimination procedure [29].
An adjustment was made for potential confounding factors, being these factors retained in the final
model [26]. Dummy variables were used for the case of categorical variables (e.g., sex), leaving one
level as reference, and all the regression coefficients from those levels of categorical variables must
be interpreted with respect to its reference level. For the estimation of possible differences between
levels of a variable, an adjusted mean comparison test was performed, using the Di Rienzo, Guzmán
and Casanoves test with a significance level of p < 0.05 [30]. A borderline significance was considered
when p-values were between 0.1 and 0.05 [31]. All the analyses were carried out using RStudio and the
statistical software R [25], plus ‘lmerTest’, ‘mgcv’, and ‘e1071’ packages.

3. Results

3.1. Effect of Nutrient Composition on Cats’ Food Preferences

Variances (i.e., eigenvalues) for the first four principal components (PC1−PC4) were greater
than one (>1) and, altogether, they explained more than 85% of the variability found in the original
variable (i.e., nutrients). These components allowed us to summarize our data into multivariate linear
regression analyses, without losing information or minimizing such loss. In particular, the values for
these components (expressed as percentages) were 34.98%, 29.70%, 13.87%, and 10.37%, respectively.
Eigenvectors (Table 1) from these four components confirm that some nutrients are correlated. Four
major nutrient groups were identified, specifically: a mineral component (Ca, P, and ash) in PC1,
a lipid component (LIP and EE) in PC2 (see Figures 1 and 2); DM and NFE in PC3; and CF in PC4.
Furthermore, the linear regression analysis between principal components and the preference of Diet
A by cats revealed a borderline significance for the effects of PC1 (Ca, P, and ash), PC2 (LIP and EE),
and PC4 (CF) over
Animals the
2019, 9, x food preferences of cats (p = 0.059, p = 0.055 and p = 0.053, respectively).
6 of 14

Figure 1. Distribution of foodofpreference

Figure 1. Distribution of kennel
food preference cats,cats,
of kennel on the
on first twotwo
the first principal
principalcomponents
components extracted
extracted
from estimate from estimate
contents of drycontents
matter of(DM),
dry matter
crude(DM), crude(CP),
protein protein (CP), fiber
crude crude (CF),
fiber (CF),
etherether
extract (EE),
extract (EE), nitrogen free extract (NFE), ash, calcium (Ca), phosphorus (P), total lipids (LIP) and
nitrogen free extract (NFE), ash, calcium (Ca), phosphorus (P), total lipids (LIP) and metabolizable
metabolizable energy (ME) by proximal chemical analysis, acid hydrolysis, and calorimetric pump
energy (ME) bydecomposition.
food proximal chemical analysis,ofacid
Space distribution diethydrolysis, and calorimetric
variables according to componentspump food
1 and 2, decomposition.
plotted as
Space distribution of diet
their eigenvectors. variables according to components 1 and 2, plotted as their eigenvectors.
Animals 2019, 9, 372 6 of 13

Table 1. Summary table of principal component analysis (PCA), indicating the importance of each
nutritional component of diets preferred by kennel cats in food preference tests, standard deviation
(SD) and the percentage of explanation of variation linked to each principal component.

Principal Component Eigenvectors

Nutritional Components 1
PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9 PC10
DM −0.04 0.29 0.66 0.24 0.32 −0.15 −0.22 0.05 −0.31 0.38
CP −0.03 0.35 −0.38 0.51 0.49 0.12 −0.08 0.04 −0.08 −0.43
CF 0.27 0.14 0.13 −0.66 0.59 0.10 0.26 0.03 0.09 −0.14
EE −0.17 0.49 −0.01 −0.25 −0.38 −0.24 0.25 0.11 −0.55 −0.30
NFE −0.20 −0.39 0.52 0.04 −0.02 0.11 −0.11 0.19 0.00 −0.68
ASH 0.47 0.19 0.15 0.03 −0.15 −0.39 −0.28 −0.57 0.21 −0.31
Ca 0.48 0.16 0.06 0.16 −0.19 −0.17 0.06 0.74 0.31 −0.01
P 0.45 0.07 0.19 0.22 −0.23 0.69 0.30 −0.20 −0.22 0.01
LIP −0.22 0.45 0.03 −0.27 −0.20 0.46 −0.57 0.08 0.29 0.00
ME −0.39 0.30 0.25 0.19 −0.05 −0.04 0.56 −0.17 0.55 0.01
Principal Component Eigenvalues
SD 1.87 1.72 1.18 1.02 0.74 0.54 0.39 0.29 0.21 0.08
% of Variance 34.98 29.70 13.87 10.37 5.45 2.89 1.49 0.83 0.43 0.00
Cumulative % 34.98 64.68 78.55 88.92 94.37 97.25 98.74 99.57 99.99 100.0
1
Principal component (PC), dry matter (DM), crude protein (CP), crude fiber (CF), ether extract (EE), nitrogen
free extract (NFE), ash, calcium (Ca), phosphorus (P), total lipids (LIP), metabolizable energy (ME), and standard
Animals 2019, 9,(SD).
deviation x 7 of 14

Figure2.2.Distribution
Figure Distributionof of
food preference
food of kennel
preference cats, on
of kennel the on
cats, firstthe
twofirst
principal components
two principal extracted
components
from estimate
extracted fromcontents
estimateofcontents
dry matter (DM),
of dry crude(DM),
matter protein (CP),protein
crude crude fiber
(CP),(CF),
crudeether
fiberextract (EE),
(CF), ether
nitrogen free extract (NFE), ash, calcium (Ca), phosphorus (P), total lipids (LIP), and metabolizable
extract (EE), nitrogen free extract (NFE), ash, calcium (Ca), phosphorus (P), total lipids (LIP), and
energy (ME) by energy
metabolizable proximal chemical
(ME) analysis,chemical
by proximal acid hydrolysis, and
analysis, calorimetric
acid pump
hydrolysis, andfood decomposition.
calorimetric pump
Space distribution of diet variables according to components 1 and 2, plotted as their
food decomposition. Space distribution of diet variables according to components 1 and 2, plottedeigenvectors and
as
differentiated by climate season (cold and hot seasons).
their eigenvectors and differentiated by climate season (cold and hot seasons).

In Table 2, both positive and negative β values are observed (PC1 = −1.42, PC2 = −1.56, and PC4
= 2.68). Meanwhile, negative correlations between the contents of ash and CF with food preference
appears (rho = −0.269, p = 0.031; rho = −0.338, p = 0.006, respectively). In the case of Ca, its contents in
the diet had a negative correlation with food preference (rho = −0.241) and a borderline significance
Animals 2019, 9, 372 7 of 13

In Table 2, both positive and negative β values are observed (PC1 = −1.42, PC2 = −1.56, and PC4
= 2.68). Meanwhile, negative correlations between the contents of ash and CF with food preference
appears (rho = −0.269, p = 0.031; rho = −0.338, p = 0.006, respectively). In the case of Ca, its contents in
the diet had a negative correlation with food preference (rho = −0.241) and a borderline significance
(p = 0.054).

Table 2. Multivariate linear regression for selected components of Principal component (PC) analysis
as they relate with food preference by kennel cats.

Variable OR 1 SE 2 p-Value
(Intercept) 74.3906 1.3737 <2 × 10−16
PC1 −1.4203 0.7403 0.0598
PC2 −1.5687 0.8034 0.0555
PC4 2.6764 1.3598 0.0537
1 Odds ratio or estimation of the impact of the component over food preference; 2 standard error.

3.2. Effect of Sex, Breed, Age, Body Weight and Season on Food Intake by Cats
Table 3 presents the fixed effects variables that were kept in the final model for food intake
and its parameter estimators. Variables that presented a significant association with cat food intake
were: body weight, sex, autumn season, and the interaction term of male sex and body weight.
Additionally, the interaction term of season and sex was kept in the model due to its potential role as a
confounding factor.

Table 3. Odds ratio and associated statistic of fixed effects included in the final intake model of
kennel cats.

Variable OR 1 SE 2 p-Value
(Intercept) 46.90 5.21 <0.001
Weight −5.49 0.82 <0.001
Sex
Male 11.76 4.72 0.014
Season
Autumn 7.86 3.43 0.022
Spring −5.31 3.32 0.110
Summer 2.66 4.02 0.508
Weight: Sex
Male −2.19 1.05 0.038
Weight: Season
Autumn −1.68 0.99 0.089
Spring 0.52 0.86 0.548
Summer 0.50 1.11 0.652
Sex: Season
Male: Autumn 4.20 2.52 0.096
Male: Spring 0.97 2.40 0.686
Male: Summer 4.18 2.67 0.119
1 Odds ratio or estimation of the impact of each variable over food intake; 2 standard error.

In the case of body weight, there was a significant effect over food intake in cats (β = −5.490;
p < 0.001). Specifically, we observed that food intake (corrected by metabolic weight) decreased by 5.490
units for every increase of one unit of body weight. Sex also had a significant effect as males increased
their food intake by 11.758 units (β = 11.758; p = 0.014). As for the season, dummy variables signaled
winter as the reference category, where autumn was significantly different than winter (β = 7.864;
p = 0.022). Thus, food intake (corrected by metabolic weight) increased by 7.864 units in autumn
vs. winter. Also, we observed a seasonal effect on food intake when reclassifying seasons simply as
either hot (summer and spring) or cold (winter and autumn), as cats ate less from Diet A during the hot
Animals 2019, 9, 372 8 of 13

season (25.84 in hot season vs. 27.96 g in cold season, SEM 0.98 g; β = −2,117; p = 0.032). Meanwhile,
males tended to decrease food intake as their body weight increased (β = −2.187; p = 0.038), whereas
the interaction between sex and season did not significantly affect the intake of Diet A (p = 0.149).
In particular, females were the only sex that significantly increased their intake under cold temperatures
Animals 2019, 9, x 9 of 14
(p = 0.002 vs. p = 0.976 in males; see Figure 3).

Hot Season Cold Season

40 *

30
Food Intake (g*BW0.75)

20

10

0
Female cats Male cats
Figure 3. Cat’s food intake means and standard errors of most consumed diets (Diet A) after two feeder
Figure 3. Cat’s food intake means and standard errors of most consumed diets (Diet A) after two
20-h preference tests according to animal’s sex (females or males) and climate season (hot or cold).
feeder 20-h preference tests according to animal’s sex (females or males) and climate season (hot or
* p < 0.05.
cold). *p < 0.05.
3.3. Effect of Sex, Breed, Age, Body Weight, and Season on Food Preferences in Cats
3.3. Effect of Sex, Breed, Age, Body Weight, and Season on Food Preferences in Cats
The mixed model analysis revealed that food preferences data was not normally distributed,
henceThedatamixed
weremodel analysis
transformed revealed
using that food
the decimal preferences
logarithmic data was
function not normally
to ensure distributed,
its normality for the
hence data were transformed using the decimal logarithmic function to ensure
residual analysis. This analysis revealed that the preference for Diet A was affected by only two its normality for the
residual
intrinsic analysis.
variables:This analysis
body weightrevealed
and sex. that
Thethefirstpreference
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and clearly
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variable,
and our analysis revealed (Table 4) that males did not prefer Diet A as much as
females (β = −8.419
Meanwhile, theand p = 0.023).
season of the year affected food preference for Diet A, as it decreased in summer
(β = −10.027 and p < 0.001) when it was compared against data for winter (via dummy variables). Also,
Table 4. Odds
the sex*season ratio anddid
interaction associated
have anstatistic of fixed
effect over effects
food included
preference forinDiet
the final
A, aspreference model of
males preferred Diet A
kennel cats.
more often during summer (β = 10.216 and p = 0.023), whereas females presented similar degree of
the year (p = 0.950).
preference for Diet A throughoutVariable OR 1 The SE interaction
2 p-Valuebetween sex and season when
food preferences were reclassified as either
(Intercept) hot (summer
76.22 2.95 and spring)
<0.001 or cold (winter and autumn) is
expressed in Figure 4. Weight −2.92 1.46 0.046
Weight^2* −0.29 0.65 0.661
Weight^3* 1.16 0.37 0.002
Sex
Male −8.42 3.60 0.023
Season
Autumn −1.59 2.31 0.493
Spring −0.76 2.25 0.736
Summer −10.03 2.72 <0.001
Sex: Season
Male: Autumn 7.62 4.32 0.078
Male: Spring 7.78 4.16 0.062
Animals 2019, 9, 372 9 of 13

Table 4. Odds ratio and associated statistic of fixed effects included in the final preference model of
kennel cats.

Variable OR 1 SE 2 p-Value
(Intercept) 76.22 2.95 <0.001
Weight −2.92 1.46 0.046
Weight 2* −0.29 0.65 0.661
Animals 2019, 9, x 10 of 14
Weight 3 * 1.16 0.37 0.002
Sex
1 Odds ratio or estimation of the impact of each variable over food preference; 2 standard error; *
Male −8.42 3.60 0.023
polynomial term of weight Season
(square and cubic terms), indicating a change in the slope at different
weights affecting food preference.
Autumn −1.59 2.31 0.493
Spring −0.76 2.25 0.736
Meanwhile, the season of the year affected
Summer food preference
−10.03 2.72 for Diet A, as it decreased in summer
<0.001
(β = −10.027 and p < 0.001)Sex:when it was compared against data for winter (via dummy variables).
Season
Also, the sex*season interaction did have an effect
Male: Autumn 7.62 over food preference0.078
4.32 for Diet A, as males preferred
Male: Spring 7.78 4.16 0.062
Diet A more often during summer (β = 10.216 and p = 0.023), whereas females presented similar
Male: Summer 10.22 4.65 0.028
degree of preference for Diet A throughout the year (p = 0.950). The interaction between sex and
1 2 standard error; * polynomial term
seasonOdds
whenratio or estimation
food of the impact
preferences of each variableas
were reclassified over food preference;
either hot (summer and spring) or cold (winter
of weight (square and cubic terms), indicating a change in the slope at different weights affecting food preference.
and autumn) is expressed in Figure 4.

Hot Season Cold Season

100

90
Food preferences (%)

*
80

70

60

50
Female cats Male cats
Figure 4. Cat’s food preferences mean and standard errors of most consumed diets (Diet A) after two
Figure 4. Cat’s food preferences mean and standard errors of most consumed diets (Diet A) after two
feeder 20-h preference tests according to animal’s sex (females or males) and climate season (hot or
feeder 20-h preference tests according to animal’s sex (females or males) and climate season (hot or
cold). * p < 0.05.
cold). *p < 0.05.
4. Discussion
4. Discussion
Cats can accept or reject some diets due to their ability of regulating their food intake according to
theirCats can accept
nutritional or reject some
requirements diets due
[4,6,9–11]. to their ability
Interestingly, of regulating
the effect their
of intrinsic andfood intake
extrinsic according
variables on
to their
their nutritional
feeding requirements
behavior has been [4,6,9–11]. Interestingly,
poorly studied theunderstanding
[5,32], but effect of intrinsic
howand extrinsic
these variables
variables may
on their feeding
influence behavior has
food preferences inbeen poorly
cats is highlystudied [5,32],
relevant for but understandingofhow
the development these
novel variables diets
commercial may
influence food preferences in cats is highly relevant for
that cater to specific populations and physiological conditions [32].the development of novel commercial diets
that cater to study,
In this specificwepopulations
assessed the and physiological
effect conditions
of nutritional [32]. on food preferences in cats and
components
found a negative correlation with mineral components (Ca, P, and on
In this study, we assessed the effect of nutritional components food
ash). In preferences
this regard,init cats and
is worth
found
notingathatnegative correlation
the content of Ca with
and Pmineral
in mostcomponents
diets used for(Ca,
theP,preference
and ash). tests
In this regard,
were it is
higher worth
than the
noting that the content of Ca and P in most diets used for the preference tests
percentages recommended by The Association of American Feed Control Officials [33], hence it is were higher than the
percentages
possible thatrecommended
cats had chosen bydiets
The Association of American
with lower contents Feedminerals
of these Controlbecause
Officialsthat
[33],might
henceavoid
it is
possible
metabolic that cats had chosen
imbalances dietsadversely
that could with lower contents
affect of these
their health minerals
[1]. becausemaintaining
For instance, that might avoid
P and
metabolic imbalances that could adversely affect their health [1]. For instance, maintaining P and Ca
within physiological levels (0.5%−0.9% and 0.6%−1.0%, respectively) in cat diets should avoid
potential health problems, such as oxalate uroliths [34] or nutritional hyperparathyroidism. In
addition, high calcium levels in cat diets can lead to depression, weakness, muscle spasms, anorexia,
polyuria, and cardiac arrhythmias due to hypercalcemia [35]. Our results also suggest that dietary
Animals 2019, 9, 372 10 of 13

Ca within physiological levels (0.5%−0.9% and 0.6%−1.0%, respectively) in cat diets should avoid
potential health problems, such as oxalate uroliths [34] or nutritional hyperparathyroidism. In addition,
high calcium levels in cat diets can lead to depression, weakness, muscle spasms, anorexia, polyuria,
and cardiac arrhythmias due to hypercalcemia [35]. Our results also suggest that dietary fiber decreases
cats’ food preferences. Although fiber content is a necessary for a regular intestinal transit in mammals,
it is negatively correlated with the nutritional content of diets intended for non-ruminants and cats
may reject high inclusions on their diets. On the other hand, though an important benefit of fiber is
its role in regulating body weight in animals that tend to overeat [36], diets that contain high levels
of soluble fibers may cause diarrhea [37], which could also result from a poor choice of fiber source,
as cats do not base their diet in the consumption of carbohydrates in large quantities [36].
Interestingly, the consumption of preferred diets was affected by body weight and sex in cats,
but not by their age. In the case of body weight, the lower consumption of Diet A (preferred diet) that
was observed in heavier animals could stem from the fact that a smaller body surface area would
generate a lower specific metabolic rate (mL of O2 g−1 h−1 ), reflecting a lower requirement of calories
per unit of weight to maintain their daily energy requirements. Another interesting feature of the data
is that heavier cats could prefer Diet A more due to some perception changes for certain macronutrients,
which may arise due to failures in the physiological regulation of food intake (e.g., leptin resistance),
as it has been reported for obese or overweight humans and rats [38,39], as well as due to a higher
tolerance threshold for high fat diets [6,18].
As for the effect of sex, it manifested as a greater consumption of Diet A by male cats and it could
relate to hormonal differences. In this regard, some studies have shown that androgens have anabolic
effects in rats and humans, which caused greater muscle mass development in males, as well as higher
metabolic rates per unit of weight than in females [40]. On the other hand, oestrogens (particularly
oestradiol) are catabolic hormones that decrease the consumption of food [38,41]. Though our results
did not reveal an effect of age on consumption of Diet A, it is plausible that an age-related sensory
deterioration could be countered by their eating behavior patterns, which is characterised by several
small rations distributed throughout the day. This would allow food to stay for a longer time within
the digestive tract and then result in an increased use of nutrients [4]. A situation that was possible to
observe in this work due to the decision to perform long-term trials that lasted for 20 hours.
As for the effect of seasons, it is well documented that environmental temperature affects food
consumption in mammals [42,43], and the thermal comfort zone for cats ranges from 15 to 25 ◦ C [44].
The increased consumption of Diet A during the cold season could be explained as an adjustment of
caloric intake that follows the environmental temperature, thus the higher the temperature, the lower
the heat loss, which decreases the consumption of the cat [45]. Results are consistent with studies
performed in Europe, which demonstrate an increase in food consumption during the cold season [46].
Though a sex*season interaction over the consumption of Diet A by cats was not observed, the season
did most markedly affect food consumption in females during hot months. This could relate to the
seasonally polyestrous nature of cats, which cyclically alternate pro-oestrus and oestrus stages during
the hot season following the increase in light hours [47]. The importance of those two stages is that they
lead to increased serum oestradiol, which in turn leads to less food consumption in females [41,47].
Some studies in rats indicated that the effect of oestradiol relates to food-intake mediators by increasing
the anorexic effect of cholecystokinin, serotonin and leptin, as well as reducing the orexigenic effect of
ghrelin [38,48,49]. However, the fact that females ate more during the cold season might be explained
by their smaller size compared to males, thus they lose heat more easily and require a higher caloric
intake to compensate for it. In this regard, some studies have found in rats and humans that females
increase their consumption of energetic diets during dioestrum [48]. And though the reproductive
cycle of cats is different than in those species by lacking a dioestrum stage, plasma concentrations
of oestradiol in queens are lower during the anoestrus stage that they undergo during the cold
season—which coincides with a short photoperiod—and such decline could then explain the increase
in food consumption [47,48].
Animals 2019, 9, 372 11 of 13

Regarding the effect of sex*season interaction on food preferences, we observed that during the
hot season only males tended to increase their preferences for Diet A. Interestingly, when Asarian and
Geary [38] worked with rats, they reported that oestradiol seems to have no effect on food preferences.
However, the increased food preferences for Diet A of males during the hot season might be explained
because males could need more exposure to volatile cues in order to better discriminate foods via their
olfactory system [13]. This is consistent with some studies in human populations that suggest lower
sensitivity thresholds to olfactory and gustatory stimuli in women as the basis for their ability to more
efficiently detect low amounts of nutrients or volatile cues [50,51]. Hence, it is likely that females cats
are also more sensitive to volatile cues and may not need higher temperatures to discriminate better
between diets that differ on their palatability. This ability to detect volatile cues in the environment
helps females avoid poisoning in complex periods such as pregnancy or lactation. Moreover, it has
been studied that preferences for flavors are affected by pre and postnatal learning processes in kittens
and other species, indicating that female flavor identification during the perinatal period could affect
the selection of a particular diet on their offspring [52].

5. Conclusions
Results suggest that body weight, sex, season, and nutritional factors may affect food preferences
in domestic cats. In particular, we found that excesses in the mineral component of the diets (Ca, P, ash),
as well as the contents of crude fiber have a negative effect on food preferences. On the contrary, body
weight decreased food consumption but increased the preference of our cats for palatable diets (Diet A).
Additionally, while males usually ate more than females, sex did not affect food preferences. As for the
effect of seasons, the fact that Diet A was preferred in the cold season was due to a higher consumption
by females, which could relate to their reproductive cycle. Yet males tended to prefer this diet more
during the hot season, probably because higher temperatures may aid them to better discriminate
palatable foods by their volatile cues. Consequently, accounting for these factors when performing
food preference tests—to a homogeneous feline population and under controlled environmental
conditions—would help to eliminate possible biases and to reduce data variability. Such results
would more reliably reveal the food preferences and feeding behavior of domestic cats when offered
commercial diets. In turn, this may improve formulation and development of cat foods adjusting to fit
animal’s specific nutrient requirements as well as to an adequate palatability level increasing both,
cats’ health and wellbeing.

Author Contributions: J.I.E., V.S., and J.F. conceived and designed the experiments; J.I.E., V.S., and J.F. performed
the experiments; R.A.A.-M. analyzed the data; R.A.A.-M., S.A.G.-P., and J.F. wrote the paper, which was revised
by all authors.
Funding: This research received no external funding.
Acknowledgments: The authors want to thank both the staff at the Research Centre for Pet Feeding Behaviour
and at the Department of Clinical Sciences from the Faculty of Veterinary and Animal Sciences of the University of
Chile for their help with missing procedural data and cat background checks, respectively. We also want to thank
Dominic Dwyer for his valuable contributions to this manuscript on the English language, writing assistance.
and proofreading the article.
Conflicts of Interest: The authors declare no conflicts of interest.

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